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A sinistral canaliculata (: ).

Article · January 1990

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Nestor J. Cazzaniga Universidad Nacional del Sur

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A SINISTRAL (GASTROPODA: AMPULLARIIDAEj1.­ Nestor J. Cazzaniga and Alejandra L. Estebenet - The genera of the family Ampullariidae show three coiling types. Montfort, from Africa, is a classical example of hyperstrophy (i.e., a sinistral coil of the shell with a dextral coil of the soft parts). Gray, from Central and South America, have dextral young (Pilsbry, 1933) but planispiral adults. The remaining genera ( Reding, Pomacea Perry, d'Orbigny, etc.) are normally dextral (Michelson, 1961). There are few known exceptions to this scheme. No dextral Lanistes have been recorded, and Dartevelle (1953) cited only four cases of individual shell sinistrality restricted to three of Pila; two shells of (Swainson) and one of Pila carinata (Swainson) from Asia (Nevill, 1877, 1886), and a single sinistral Pila wernei dewulfi Bequaert & Clench from Africa, all of which were found empty. Additional recorded cases of sinistrality in this genus are for Pila conica (Wood) (Letson, 1897), and (Linnaeus) (Walker, 1917). A live sinistral specimen of Pila pesmei (Morelet) was collected in Thailand in 1979 (R. Keawjam, pers. comm.). Emery (1943) reported a sinistral (Say) from Florida, U.s.A., and, in an ecological study, Burky (1974) mentioned three sinistral spat and one sinistral juvenile Pomacea urceus (Muller) from Venezuela. These are the only cases of sinistrality in the genus Pomacea known to us. A live sinistral female was found in a stock of ca. 350 dextral Pomacea canaliculata (Lamarck, 1822) in Paseo del Bosque (La Plata City, Argentina). It was maintained in a 10 liter with two dextral males of similar size. Matings were observed. on January 8 (9:30 PM) and January 13 (9:00 AM) 1989, but no spawning occurred. On January 18 both of these dextral males were replaced by similar dextral ones, but the female still failed. to spawn and eventually died on January 27. It was therefore not possible to trace the genetic origin of the sinistrality. The behaviour of the sinistral was normal. It actively fed on lettuce. The protrusion of the aerial on the right confirmed that it was a case of visceral inversion and not an individual hyperstrophy. This fact would explain the failure of oviposition, since the penis sheath was observed to enter the pallial chamber of the female via the usual right side or slightly more towards the center, whereas the vagina was on the left. Morphology. The shell (Fig. 1) corresponds to the general features of the species, even in the shape of the last whorl, which is rather square. It shows a smooth "martelee" sculpture. The umbilicus is wide. The periostracum is thin, light brown, and bears 21 coloured spiral stripes, which are also evident on the inside. The spire is not eroded.. The shell is 46.5 mm long and 41.0 mm wide. The last whorl is 41.9 mm long (90.1 % of the total length). The specimen has 5 3/4 whorls. The aperture measures 33.3 x 24.5 mm. The is a mirror-image but otherwise identical to the dextral one and measures 29.5 x 18.1 mm; the width/length ratio of the operculum (61.4%)lies within the normal range for female shells of this species (Cazzantga, in press). Fig. 2 shows the general anatomy of the cephalopodium and pallial chamber of the sinistral Pomacea canaliculata. In this specimen it is the right nuchal lobe which has transformed into a siphon. All the organs are well developed but inversely disposed. The rudimentary penial complex (Andrews, 1964) is represented here by a triangular fold of the left edge of the , without the small penis drawn by Hylton Scott (1957). The specimen was fully mature, with a large albumen gland of normal shape and a rather inflated pallial oviduct. Location of the material. The shell, operculum and dissected soft parts have been deposited in the Museo Argentino de Ciencias Naturales "Bernardino Rivadavia"

1ContTibution No. 28 of the Laboratorio de Ecologfa Acuatica, Departamento de Biologta de la Universidad Nacional del Sur. The first author (NJC) is a researcher of the Comisi6n de Investigaciones Cientfficas de la Provincia de Buenos Aires (Argentina). (99) 100 Malacological Review, Vol. 23, 1990

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RG. 1. Pomacea canaliculata. A dextral and the sinistral shell and operculum.

os PF

FIG. 2. General aspect of the anatomy of the cephalopodium and pallial chamber in the sinistral Pomacea canaliculata. AG: albumen gland; AP: anal papilla; CT: ctenidium; E: eye; F: foot; LNL: left nuchal lobe; OS: osphradium; PF: pallial fold ; PO: pallial oviduct; RNL: right nuchal lobe; 5: snout; T: tentacle; VP: vestigial penial complex. Brief Communication - Cazzaniga and Estebenet 101

(Buenos Aires), under the number 32,950 of the Mollusc Collection. Remarks. Sinistrality in normally dextral Ampullariidae is an infrequent phenomenon. Paseo del Bosque pond is situated about 100 m from the Museo de La Plata, and has long been used as a source of material for malacological research and university courses on invertebrates. Hylton Scott (1934, 1957) collected Pomacea canaliculata there for her embryological and anatomical studies. The senior author has worked with this species since 1978, frequently obtaining specimens from the same site (Cazzaniga, 1981, 1983). Martin (1980) carried out a study on the penial complex of this species, including samples from Paseo del Bosque. However, in all previous collections of Pomacea from Paseo del Bosque pond, this is the first known sinistral specimen. This accords with the fact that there are very few antecedents of sinistrality within the family as a whole. Milward de Andrade et al. (1978) and Guimaraes (1981) carried out extensive field observations on the closely related species (Reeve) from Brazil, but made no mention of sinistral specimens. Bourne (1983), Snyder & Kale (1983) and other authors have measured several thousand Pomacea shells from feeding deposits of the (Rostrhamus sociabilis, Aves, Accipitridae), and no sinistral shells were reported. Sinistral specimens of the Asian and African Pila have been reported rarely, with a total of only six shells reported in over a century. Burky's (1974) sinistral specimens of Pomacea urceus were newly hatched and a juvenile. No adult sinistral shells were cited, though P. urceus is a well known species, repeatedly cited from northern South America (Baker, 1930; Pain, 1950, 1956, 1960). Sinistrality in normally dextral species seems to be more frequent, however, in pulmonate shells. No mechanical barrier appears to exist for copulation between dextral and sinistrallymnaeas (Boycott et al., 1930), and other gastropods. Nevertheless, Lipton & Murray (1979) stated that some degree of assortative mating appears in mating populations of Partula spp., which should enforce reproductive isolation of dextral and sinistral . The left coiled shells of Pila and Pomacea must be considered developmental accidents or individual mutations. Should the sinistrality be due to the former reason, then the cause of the condition has no genetic basis. Should the sinistrality be due to the latter reason, then the mutated gene is destined to disappear when it occurs in single specimens, since normal copulation is impeded by mechanical incompatibility. We wish to thank Drs. R. Keawjam ana J. B. Burch for providing some information and bibliography, and for reading the manuscript. LITERATURE CITED. - ANDREWS, E.B. 1%4. The functional anatomy and histology of the reproductive system of some pilid gastropod molluscs. Proceedings of the malacological Society of London, 36: 121-140. - BAKER, H.B. 1930. The collected by the

University of Michigan w Williamson expedition in Venezuela. Occasional Papers of the Museum of Zoology, University of Michigan, (210): 1-95. - BOURNE, G.R. 1983. Snail Kite feeding ecology: Some correlates and tests of optimal foraging. Doctoral Dissertation, University of Michigan, Ann Arbor. xi + 110 pp. -- BOYCOTT, A.E., DIVER, C, GARSTANG, S.L., (Mrs. A.C Hardy), & TURNER, F.M. 1930. The inheritance of sinistrality in Lymnaea peregra (Mollusca: ). Philosophical Transactions of the Royal Society of London, B, 219: 51-131. -- BURKY, A.j. 1974. Growth and biomass . production of an amphibious snail, Pomacea urceus (Muller), from the Venezuelan savannah. Proceedings of the mJllacological Society of London, 41: 127-143. - CAZZANIGA, N.j. 1981. Evaluaci6n preliminar de un gaster6podo para el control de malezas acuaticas sumergidas. Pp. 131-165 In: Reuni6n sobre Malezas Subacuaticas en Canales de Desagiie de CORFO, 2, Pedro Luro, February 1981. La Plata. -- CAZZANIGA, N.j. 1983. Apple snails eating Chura, Aquaphyte, 3(2): 1, 4. -- CAZZANIGA, N.j. in press. Sexual dimorphism in Pomacea canaliculata (Gastropoda: Ampullariidae). The Veliger. -­ DARTEVELLE, E. 1953. A sinistral Pila from Africa. Journal ofConclwlogy, 23(9): 288-289, pI. 8. - EMERY, D.L. 1943. Sinistral Pomacea. The , 57(2): 66. -- GUIMARAES, CT. 1981. Algumas observacoes de campo sobre biologia e ecologia de Pomacea haustrum (Reeve, 1856). (Mollusca: Pilidae), Memorias do Instituto Oswaldo Cruz, 76(4): 343-351. ­ HYLTON SCOTT, M.l. 1934. Sobre el desarrollo embrionario de canaliculata. Revista del Museo de La Plata, 34: 373-385. -- HYLTON SCOTT, M.l. 1957. Estudio 102 Malacological Review, Vol. 23, 1990

morfo16gico y taxon6mico de los ampullaridos de la Republica Argentina. Revista del Museo Argentino de Ciencias Naturales "Bernardino Rioadaoia", Ciencias ZooI6gicas, 3(5): 233­ 333. - LETSON, Jennie E. 1897. Sinistral Ampullaria. The Nautilus, 11(3): 33-34. ­ LIPTON, CS. & MURRAY, J. 1979. Courtship of land snails of the genus Partula, Malacologia, 19(1): 129-146. -- MARTIN, S.M. 1980. Contribuci6n al estudio histol6gico del complejo peniano de algunes ampularias argentinas (Mollusca: Ampullariidae). Neotropica, 26(75): 47-52. - MICHELSON, E.H. 1961. On the generic limits in the family Pilidae (Prosobranchia: Mollusca). Breoiora, Museum of Comparative Zoology, Harvard University, 133: 1-10. -- MILWARD DE ANDRADE, R., CARVALHO, O. dos 5. & GUlMARAES, CT. 1978. Alguns dados bioecol6gicos de Pomacea haustrum (Reeve, 1856), predador-competidor de hospedeiros intermediarios de Schistosoma mansoni Sam ban, 1907. Revista de SaUde Publica, 12: 78-89. -- NEVILL, Geoffrey. 1877. Catalogue of Mollusca in the Indian Museum, Calcutta, fasc, E, p. 3. - NEVILL, Geoffrey. 1886. Hand List of Mollusca in the Indian Museum, Calcutta, pt. 2, pp. i-x, 1-306. -- PAIN, T. 1950. Pomacea (Ampullariidae) of British Guiana. Proceedings Ofthe malacological Society at London, 28(2-3): 63-74, pI. 6-8. - PAIN, T. 1956. On a collection of Pomacea from Colombia, with description of a new subspecies. Journal of Conchology, 24(3): 73-79. - PAIN, T. 1960. Pomacea (Ampullariidae) of the System. Journal of Conchology, 24(12): 421­ 432. - PIL5BRY, H.A. 1933. Zoological results of the Matto Grosso Expedition to Brazil in 1931. II. Mollusca. Proceedings of the Academy of Natural Sciences of Philadelphia, 85: 67-76. ­ - SNYDER, N.R.F. & KALE, K.W., II. 1983. Mollusk by snail kites in Colombia. The Auk, 100: 93-97. WALKER, Bryant. 1917. A sinistral Ampul/aria. The Nautilus, 31(1): 35-36. • • • • •

NESTOR J. CAZZANIGA and ALEJANDRA L. ESTEBENET, Laooratorio de EcologCa Acu4tica, Departamento de Biologia, Universidad Nacional del Sur, 8000 Bahfa Blanca, Argentina.

Ma1Qcological Review, P. O. Box3037, Ann Arbor, Michigan 48106, U.S.A. e socery for Experimental and Descriptive Malacology, 1990

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